Neogene lake systems of Central and South-Eastern Europe ...

Abstract 

Neogene lake systems of Central and South-Eastern Europe: 

Faunal diversity, gradients and interrelations 

Mathias Harzhauser ⁎ , Oleg Mandic 

Naturhistorisches Museum Wien, Geologisch-Paläontologische Abteilung, Burgring 7, A-1010 Wien, Austria 

Received 19 September 2007; received in revised form 10 December 2007; accepted 18 December 2007 

The gastropod γ-diversity of 12 Neogene lake systems is evaluated. In total, 1184 gastropod taxa from 119 localities are recorded deriving 

from the Early Miocene Rzehakia Lake System, the Early to Middle Miocene Dinarid Lake System, Lake Skopje, the Paratethyan Sarmatian lakes 

and the South German lakes, the Late Miocene Lake Pannon, the Pliocene lakes Dacia, Transylvania, Slavonia, Kosovo and Šoštanj as well as 

the Holocene Lake Petea. Each lake system is characterised according to its faunistic inventory and endemism. According to their gastropod 

faunas the lakes may be divided into pyrgulid-, hydrobiid-, viviparid- and planorbid-dominated ones. The generally high endemism rate is 

between 60 and 98%. Species diversity and generic diversity are strongly correlated. In contrast, neither endemism nor lake size are tightly linked 

with γ-diversity. Outstandingly high diversities such as observed for Lake Pannon are rather a result of the combined effect of autochthonous 

evolution in a long-lived system and accumulation of inherited elements. Examples of parallel evolution in lymnaeids and planorbids are presented. 

© 2008 Elsevier B.V. All rights reserved. 

Keywords: Gastropods; Freshwater molluscs; Endemism; Ancient lakes; Evolution 

1. Introduction 

The Central and South-Eastern European freshwater and 

brackish systems of the Neogene are often characterised by 

outstanding endemisms. Despite the enormous amount of systematic 

papers dealing with local faunas, the relations between 

these lake systems in space and time are still unexplored. The 

most important biogeographic entities are the Early Miocene 

Rzehakia Lake System, the Early to Middle Miocene Dinarid 

Lake System, Lake Skopje, the Paratethyan Sarmatian lakes and 

the South German lakes, the Late Miocene Lake Pannon, the 

Pliocene lakes Dacia, Transylvania, Slavonia and Kosovo, as 

well as the Pleistocene and Holocene lakes Šoštanj and Petea 

(Fig. 1). The main obstacle for taxonomists working with the 

Neogene lake faunas of Central and South-Eastern Europe is the 

complex paleogeographic situation. Some areas have been 

repeatedly covered by different lake systems and therefore a 

⁎ Corresponding author. Fax: +43 1 52177 459. 

E-mail address: mathias.harzhauser@nhm-wien.ac.at (M. Harzhauser). 

0031-0182/$ - see front matter © 2008 Elsevier B.V. All rights reserved. 

doi:10.1016/j.palaeo.2007.12.013 

Available online at www.sciencedirect.com 

Palaeogeography, Palaeoclimatology, Palaeoecology 260 (2008) 417–434 

single literature-based locality name might represent completely 

different faunas. Moreover, several classical monographs 

intermingled faunas from several lake systems and very 

different stratigraphic levels (e.g. Neumayr 1869, 1880; Brusina 

1897, 1902a). Especially, the separation of Lake Pannon faunas 

from those of the older Dinarid Lake System was completely 

obscure for most taxonomists (e.g. Nuttall, 1990). Molluscs of 

some ancient Balkan lakes such as the Early Miocene Lake 

Sumadija or the early Middle Miocene Lake Serbia (Krstić 

et al., 2001, 2003, 2007) are still insufficiently documented and 

therefore excluded from present analysis. 

2. Methods and limitations 

www.elsevier.com/locate/palaeo 

The dataset is based on a compilation of published gastropod 

faunas from Central and South-Eastern European Neogene lake 

systems. The results of more than 120 systematic papers have 

been integrated; details and references are given below in 

section 4. In addition, material from the collection of the Natural 

History Museum Vienna (NHM) has been included. The full

418 M. Harzhauser, O. Mandic / Palaeogeography, Palaeoclimatology, Palaeoecology 260 (2008) 417–434

M. Harzhauser, O. Mandic / Palaeogeography, Palaeoclimatology, Palaeoecology 260 (2008) 417–434 

dataset with detailed locality information is available online: 

http://www.nhm-wien.ac.at/Content.Node/forschung/geologie/ 

mitarbeiter/pdfs/Harzhauser-Mandic-freshwater.xls. The 

recorded species-level taxa have been systematically arranged 

and listed according to localities. In a next step, localities have 

been grouped into geographic and stratigraphic units. Hierarchical 

cluster analysis and non-metric multidimensional 

scaling (nMDS) were performed using the statistic software 

packages PAST (Hammer et al., 2001) and PRIMER (Clarke 

and Warwick, 1994). The groupings achieved by means of 

hierarchical cluster analysis were tested for robustness by using 

different available algorithms and were additionally compared to 

results from nMDS. The similarity measures providing the best 

interpretable groupings were the Euclidean distance (between 

rows) for the percentage contribution data and the Simpson 

index and Bray–Curtis measure for the presence/absence data. 

Generally, we follow the systematic groupings of Wenz 

(1923–1930, 1938–1944), Falkner et al. (2001), Bank et al. 

(2001) and Harzhauser et al. (2002). Nevertheless, the herein 

presented affiliation of several genera (e.g. Staja Brusina, 1897, 

Bania Brusina, 1896, Gyromelania Wenz, 1938–1944, Scalimelania 

Wenz, 1938–1944) to higher taxa is problematic and 

may change after a modern revision. Moreover, it has to be kept 

in mind that the number of described species in the literature is 

too high for many genera as modern revisions are missing for 

most groups. A taxonomic revision, however, is beyond the 

scope of this study. Despite these limitations and “taxonomic 

noise”, we think that this dataset is a serious first approximation. 

3. The lakes: geography, geological settings and 

stratigraphic framework 

3.1. Rzehakia Lake System (RLS, ~17.5–17.2 Ma; S. Germany, 

Austria, Moravia) 

The oldest lake system treated herein is the Early Miocene 

Rzehakia Lake System (Fig. 2). Its name is derived from the 

endemic bivalve genus Rzehakia (Korobkov, 1954). The 

geographic extension of the RLS reaches from Bavaria to 

Moravia, covering an area about 650 km long (W–E) and 

150 km wide (N–S) (Senes, 1973). It had formed along the 

northern shoreline of the Paratethys Sea in the North-Alpine 

Foreland Basin and, in its Moravian prolongation, in the 

Carpathian Foredeep. The lakes developed during the Early 

Miocene (mid-Burdigalian) and reflect the sea-level lowstand 

TB 2.1 of Haq et al. (1988) (Rögl, 1998). This event caused the 

Central Paratethys Sea to disintegrate into several basins and 

allowed the development of strongly structured coastal plains 

with extensive brackish-water lakes. Little is known about the 

geochemistry of these lakes. Traditionally, they are considered 

as marine-derived brackish lakes (Rögl, 1998; Popov et al., 

2004). This assumption is supported by the occurrence of 

endemic cardiids such as Limnopagetia (Schlickum, 1963) and 

Limnopappia (Schlickum, 1962) and taxa such as Siliqua 

(Megerle von Mühlfeld, 1811), which all have marine ancestors. 

The origin of the RLS fauna is partly rooted in the Eastern 

Paratethys (Popov et al., 1993; Rögl, 1998; 1999). Several 

endemic genera apparently originated in the faunas of the 

Kozakhurian Stage (e.g. Limnopagetia) and subsequently 

settled the western and central Paratethyan shores. 

Based on considerable differences of the mollusc faunas of the 

western (Bavarian) and eastern (Austrian–Moravian) part, 

Harzhauser and Mandic (2008) and Mandic and Corić (2007) 

proposed the existence of at least two disconnected lakes. A 

further separation into smaller lakes in the western part of the RLS 

is indicated by slight faunistic differences between the Lower 

Bavarian Oncophora Basin and the Upper Bavarian Kirchberg 

Basin (Kowalke and Reichenbacher, 2005). The paleobiogeographic 

relations within the RLS are based on the high degree of 

endemics on the generic level. All lakes of that system have in 

common taxa such as the bivalves Rzehakia, Limnopagetia, and 

Limnopappia and the gastropod Ctyrokia (Schlickum, 1965). 

On the species-level, however, hardly any faunistic relation is 

represented aside from the ubiquist Melanopsis impressa (Krauss, 

1852) along with the theodoxid Theodoxus cyrtocelis (Krauss, 

1852) and the planorbid Gyraulus applantus (Thomae, 1845). 

The absence of RLS bivalves in the probably partly synchronous 

Dinaride Lake System suggests a distinct paleobiogeographic 

boundary between these lake systems. 

3.2. Dinarid Lake System (DLS, ~17–15 Ma; Croatia, Bosnia 

and Herzegovina, Montenegro, Serbia, Hungary and Slovenia) 

The Dinarid Lake System (DLS) formed during the late 

Oligocene and Miocene in today NW–SE trending intramountain 

basins parallel to the slowly rising Dinarid mountain chains 

(Pavelić, 2001). Extensional tectonics generated enhanced 

subsidence of elongated depressions during the Early to Late 

Miocene. The comparatively low terrigeneous input supported 

the diversification of lacustrine environments, including both 

deep- and shallow-water habitats. This habitat diversification 

sparked the spectacular Miocene radiation of the benthic fauna. 

Geographically, the deposits of the DLS cover parts of Croatia, 

Bosnia and Herzegovina, Montenegro, Serbia, Hungary and 

Slovenia (Krstić et al., 2001, 2003). During its maximum extent, 

the lake system covered an area of c. 75,000 km 2 . Subsequent 

rifting in the Pannonian Basin System triggered the marine 

flooding of the northern DLS and considerably reduced it. 

The stratigraphic correlation and paleogeographic extension 

of that shrunken DLS, termed Lake Herzegovina by Vujnović 

et al. (2000), are still under discussion. The younger deposits, 

however, lack DLS endemics such as Clivunella Katzer, 1918 

Fig. 1. Geographic setting of the investigated lake systems. 1: Rzehakia Lake System (RLS, ~17.5–17.2 Ma), 2: Dinarid Lake System (DLS, ~17–15 Ma), 3: Lake 

Skopje (LSK, ~15 Ma), 4: Paratethyan Sarmatian Lakes (PSL, 12.4–11.8 Ma), 5: South German lakes (incl. Lake Steinheim) (SGL, 14.3– ~12 Ma), 6: Lake Pannon 

(LP, 11.6–5.8 Ma) 7: Lake Slavonia (LS, ~4–3 Ma), 8: Lake Transylvania (LT, ~4.5–3 Ma), 9: Lake Dacia (LD, ~5–3 Ma), 10: Lake Kosovo (LK, ~3–2 Ma), 

11: Lake Šoštanj (LSO, 2.5 Ma), 12: Lake Petea (P, 0.1–0 Ma). The outlines of the Early and Middle Miocene lakes reflect only the modern sediment distribution, 

whilst the outlines of the Late Miocene and Pliocene lakes roughly coincide with their former extent. 

419



and Delmiella Kochansky-Devidé and Slišković, 1972 

(Kochansky-Devidé and Slišković, 1978, 1980). A typical 

fauna of the late DLS is recorded from the Sinj Basin in southeastern 

Croatia, It represents the best investigated record of the 

DLS and yields an extraordinarily high species diversity (e.g. 

Neumayr, 1869; Brusina, 1874, 1897, 1902a). 

3.3. Lake Skopje (LSK, ~15 Ma; Macedonia) 

The data on that lake are extremely poor. The investigated 

mollusc fauna was first described more than a century ago 

(Burgerstein, 1877; Pavlovic, 1903) and is known so far only 

from Skopje. Following the current paleogeographic reconstructions, 

those deposits could represent the southern part of 

the Serbian Lake of Krstić et al. (2001, 2003, 2007) dated 

as early Middle Miocene (c. 16–14 Ma). The Serbian Lake 

extended via a 200 km wide, NNW–SSE striking depression 

between the Dinaride and Carpathian orogenes from Belgrade 

(N Serbia) to Serres (N Greece). Its molluscs, including Kosovia 

(Pavlovic, 1903), are insufficiently documented — mainly in 

unpublished reports and theses. Apparently, based on a synopsis 

by Krstić et al. (2007), they differ not only from synchronous 

DLS faunas but also from those of Lake Skopje. Consequently, 

Lake Skopje is treated herein as an independent paleogeographic 

unit. 

3.4. Paratethyan Sarmatian Lakes (PSL, 12.4–11.8 Ma; Romania, 

Austria, Hungary) 

During the late Middle Miocene, the isolated Paratethys 

Sea developed a conspicuous endemic marine mollusc fauna 

(Harzhauser and Piller, 2007). The coastal flats of this sea were 

fringed by several freshwater systems. The paleogeography of 

these coastal lakes is unclear and our knowledge on the fauna 

might be incomplete. Only few localities yield rich assemblages, 

whereas most other occurrences are rather out-of-habitat 

findings in marine deposits. Important Sarmatian wetland faunas 

are known from Soceni in Romania (Jekelius, 1944) and the 

Austrian Eisenstadt-Sopron Basin (Harzhauser and Kowalke 

2002). Răcăştie (formerly Rákosd) in the Deva region in 

Romania is another important Sarmatian locality described by 

Gaál (1911) and Szalai (1928). In addition, Szalai (1928) and 

Boda (1959) described species from various Hungarian localities 

in the Bakony region and the Bükk Mountains. 

3.5. South German lakes (incl. Lake Steinheim) (SGL, 14.3 to 

~12 Ma; Germany) 

During the late Middle Miocene several freshwater systems 

developed in southern Germany in the area between Munich, 

Nürnberg and Stuttgart. The most important of these are the 

Steinheim Lake in the Swabian Alb and the Ries Lake at the 

border between the Swabian and Franconian Alb. Both formed 

by a simultaneous meteorite impact during the middle Miocene 

(~14.3 Ma; Tütken et al., 2006). The crater basins became filled 

by freshwater, and long-lived lakes became established. The 

smaller Steinheim Lake had a diameter of c. 3.5 km, whilst the 

Nördliger Ries impact structure was c. 25 km in diameter. Aside 

from endemics, the mollusc fauna is closely related to the 

assemblages of the coeval wetland faunas of the so-called 

Silvana Beds in the adjacent North-Alpine Foreland Basin. 

Representative localities are Hohenememmingen (26 km SSE 

of Nördlingen) and Zwiefaltendorf (70 km SE of Steinheim). 

Little is known about the paleogeography and paleolimnology 

of the associated small lakes. Especially at Zwiefaltendorf, the 

mollusc fauna was described from reworked lithoclasts outcropping 

in Pleistocene deposits (Schlickum, 1976). Despite the 

earlier impact age of 14.3 Ma, most of the assemblages are 

correlated with the mammal zone MN 7 (Tütken et al., 2006), 

pointing to an age between c. 13.5–12 Ma. 

3.6. Lake Pannon (LP, 11.6–5.8 Ma; Austria, Czech Republic, 

Slovakia, Hungary, Romania, Croatia, Slovenia, Bosnia, Serbia) 

At about 11.6 Ma a glacioeustatic sea-level drop caused 

the final disintegration of the Paratethys Sea, and Lake 

Pannon arose in the Pannonian basin system (Magyar et al., 

1999; Harzhauser et al., 2004). The benthic ecosystem 

collapsed at that point and marine life completely vanished. 

The lake was initially brackish, slowly freshening and slightly 

alkaline (Harzhauser et al., 2007). A very detailed paleogeographic 

development throughout the late Miocene is provided 

by Magyar et al. (1999). Accordingly, Lake Pannon attained a 

maximum length of 860 km (from the Karlovac Basin close to 

Zagreb in the west to the Transylvanian Basin in Romania in the 

east) and a width of 550 km (from the Vienna Basin in the north 

to Belgrade in the south). It covered an area of c. 290,000 km 2 

and is the largest aquatic system considered in this study. The 

lake was highly structured by numerous islands and mountain 

ranges. Its maximum water depth may have reached 800 m in 

its central part but less than 200 m elsewhere (Magyar et al., 

1999). 

At around 9 Ma the lake began to shrink. Its north-western 

part turned into fluvial plains and, in the east, the Transylvanian 

Basin became dry land, reducing the area to c. 180,000 km 2 . 

Finally, in the latest Miocene, a comparably small lake of c. 

480 km width remained, covering only the southern basins of the 

Pannonian basin system. Herein, the succession is separated into 

3 time slices: LP, Phase I: 11.6–10.0 Ma, Phase II: 10.0–8.0 Ma, 

Phase III: 8.0–5.8 Ma. These units roughly represent the buildup 

phase of Lake Pannon (Lower Pannonian), its maximum 

extent (Middle Pannonian, C. subglobosae Zone) and its 

gradual retreat (Upper Pannonian=“Pontian” sensu Stevanović, 

1990a,b,c). 

Fig. 2. Chronostratigraphic and biostratigraphic correlation scheme for the Mediterranean and Paratethyan areas (after Gradstein et al., 2004 and Piller et al., 2007). The 

stratigraphic range and the number of documented species and genera of each lake system are indicated in the right column (see text for abbreviations). Numbers in 

circles represent species that occur in two lakes. 

421



3.7. Lake Dacia (LD, ~5–3 Ma; Romania, Bulgaria) 

Lake Dacia formed in Pliocene times in place of the former 

Eastern Paratethys Sea. It filled the name-giving Dacian Basin 

and had a W–E extension of roughly 500 km and less than 

200 km width, covering an area of c. 78,000 km 2 . It was 

delimited in the north and west by the Carpathian Mountains 

and by the Balkanids and the Moesian platform in the south and 

extended into the area of the modern Black Sea. The age of the 

herein considered deposits is Dacian to Romanian (Snel et al., 

2006). 

3.8. Lake Transylvania (LT, ~4.5–3 Ma; Romania) 

This elongate U-shaped lake was situated on the SE- 

Carpathians and covered the Brasov Basin complex, the Ciuc 

Basin and the Gheorgheni Graben (see Fielitz and Seghedi, 2005 

for tectonic setting). It had a length of about 180 km (N–S) and 

was rather narrow, attaining a maximum width of 20–30 km. This 

only c. 4500 km 2 large lake was not a relic of Lake Pannon, but 

formed independently during the late Dacian and Romanian 

(László, 2005) within the Carpathian nappe system. 

3.9. Lake Slavonia (LS, ~4–3 Ma; Croatia, Bosnia, Serbia, 

Romania) 

This lake is also known as Paludina Lake, referring to 

the conspicuous diversity and endemic evolution of viviparid 

gastropods. It's a small Pliocene lake about 290 km long and 

120 km wide, covering the southernmost basins of the 

Pannonian basin system over about 28,000 km 2 . Its geographic 

extension is similar to that of the latest phase of Lake Pannon. 

Therefore, it is discussed by some authors as being a direct 

descendent of Lake Pannon (e.g. Magyar et al., 1999). This 

relation remains somewhat questionable because reliable geological 

data and modern datings are missing. 

3.10. Lake Kosovo (LK, ~3–2 Ma; Serbia, Kosovo) 

Lake Kosovo was a roughly circular lake of about 50 km 

diameter covering an area of c. 8000 km 2 . Its deposits are 

restricted to the larger Metohia Basin in the west and the 

elongated Kosovo Basin in the east (Atanacković, 1990). The 

stratigraphy of the basin is described by Milosević (1966). 

Accordingly, mollusc faunas occur in the Kosovo Basin in 

Middle Miocene, Upper Miocene and Pliocene deposits. The 

oldest fauna is part of Lake Serbia and is not considered herein. 

The Miocene and Pliocene faunas have been revised by Krstić 

et al. (2001), who treat Lake Kosovo as part of the Macedonian- 

Drim System (Marović et al., 1999). The latter lake system 

covered southern Serbia, Macedonia, southern Bulgaria and 

central Greece during the Late Pliocene. Its endemic mollusc 

fauna of Akchagylian age, however, is known so far only from 

the Metohia and Kosovo Basins (Atanacković, 1990; Krstić 

et al., 2001). 

3.11. Lake Šoštanj (LSO, 2.5 Ma; Slovenia) 

The investigated fauna derives from Pliocene lignite-bearing 

lacustrine deposits at Šoštanj near Velenje in Slovenia. 

Geologically, these deposits are part of the Velenje Basin and 

are dated as Villafranchian (Brezigar et al., 1985). Three small 

lakes are still present in the depression along a length of c. 4 km; 

the extent of the Pliocene lake is unknown. The locality is also 

referred to as Schönstein in the old literature (Rolle, 1860, 

1861). 

3.12. Lake Petea (P, 0.1–0 Ma; Romania) 

Lake Petea, situated about 9 km SE of Oradea in W. Romania, 

is the only still existing lake in the herein utilised dataset. It is a 

very small (N1 km 2 ) thermal-spring-fed freshwater rivulet and 

lake with constant water temperature of c. 30 °C. The area is now 

protected because of the occurrence of the endemic water lily 

Nymphaea lotus thermalis (de Candolle, 1821), the rudd Scardinus 

erithrophtalmus racovitzai (Müller, 1958) and the gastropod 

Melanopsis parreyssi (Pauca, 1933). Lake Petea is a very young, 

mainly Holocene, aquatic system which did not originate before 

Pleistocene times (pers. com. Marton Venczel). Due to the 

complex history of the region, the locality is referred to in the 

literature also as Bischofsbad (German) and as Püspökfürdö 

(Hungarian). It is included herein because of its diversity of 

melanopsids and its Mio-Pliocene “flair”, whilst other Pleistocene 

and Holocene lakes are excluded. 

4. Results 

In total, 1184 gastropod species and (chrono- or morpho-) 

subspecies from 119 localities have been compiled from 

the extensive literature on Miocene to Pleistocene Central and 

South European lakes and lake systems. This surprisingly high 

diversity is even comparable with marine gastropod diversity 

for the same geographic area during the Miocene (c. 1300 taxa 

in Harzhauser and Piller, 2007). In a first step a cluster analysis 

was performed to group localities in evolutionary entities. The 

number of taxa in single localities differed strongly, ranging 

Fig. 3. Plots of cluster analysis using Bray–Curtis similarity measure and Group Average method and non-metric Multidimensional Scaling (MDS) of Miocene to 

Pliocene lake-fauna bearing localities with more than 23 taxa. The cluster analysis demonstrates the presence of 7 taxonomically constrained paleogeographic, 

paleobiogeographic and stratigraphic units (Groups 1 to 7). The peri-Paratethys units (Groups 1 to 3) are strictly divided from units of the Central Paratethys/Lake 

Pannon region, forming a coherent but internally hierarchically ordered group. The hierarchical ordering of localities of that latter group follows a stratigraphic pattern, 

which is also evident in the non-metric MDS plot. The hierarchical/stratigraphical ordinance starts with the Middle Miocene Sarmatian Lakes (Group 4), goes through 

early Late Miocene (Group 5) and late Late Miocene (Group 6) Lake Pannon up to the latest Miocene and Pliocene residual lakes (Group 7). The subgroups within 

Group 7 reflect paleogeographic units. The full dataset with all localities is available online: http://www.nhm-wien.ac.at/Content.Node/forschung/geologie/mitarbeiter/ 

pdfs/Harzhauser-Mandic-freshwater.xls. 

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424 M. Harzhauser, O. Mandic / Palaeogeography, Palaeoclimatology, Palaeoecology 260 (2008) 417–434 

from one to maximally 97 recorded species. Prior to analysis the 

dataset was therefore filtered to records containing sufficient 

taxa for a reasonable comparison. The best results were 

achieved from the species diversity of 24 upwards and using 

the Bray Curtis Similarity measure. The resulting grouping was 

used as test for a priori assumptions about paleogeographic 

units. 

The ordering of localities coincided with their paleogeographic 

and stratigraphic patterns. Thus, seven paleobiogeographic 

units were clearly distinguished (Fig. 3): the Dinaride 

Lake System, the Paratethyan Sarmatian Lakes, the South 

German Lakes, Lake Kosovo, Lake Pannon I & II and Lake 

Pannon III, and a cluster representing the faunas of the latest 

Lake Pannon, Lake Slavonia, Lake Dacia and Lake Transylvania. 

Within the area of the former Paratethys Sea, the hierarchical 

ordering showed a clear stratigraphic pattern, with the 

oldest lake systems at the base and the Pliocene lakes at the top 

of the line-up. The peri-Paratethys systems (Dinaride Lake 

System, South German Lakes and Lake Kosovo), although 

principally following a similar stratigraphic pattern, ordered 

strictly separately from the Central Paratethys line-up, underlining 

their autochthonous evolutionary and paleoecological 

status. 

4.1. Faunistic inventories and sources 

The groupings in Fig. 3 represent ancient lake systems with 

fairly consistent faunas. In the next step the faunas were merged 

for each lake to provide a synopsis of the faunistic composition 

of each lake (Fig. 4). This dataset allows estimation of the 

γ-diversities (sensu Whittaker, 1972) and outlines the dominant 

taxa. There are, however, clear limitations to this approach. 

Especially the phenotypic plasticity of several freshwater 

gastropod species (e.g. within Melanopsis Férussac, 1823) is 

difficult to handle and might result in over-splitting in some 

genera (Geary 1990). Hybridisation effects as discussed by 

Geary (1992) and Bandel (2000) for Lake Pannon melanopsids 

will also increase the inventory. As this phenomenon cannot be 

solved based solely on conchological data, we maintain several 

of these critical morpho-species. Another drawback is the timeaveraging 

that is inevitable in such datasets. Therefore, at least 

for the extremely long-lived Lake Pannon, we tried to separate 

the faunas into 3 time slices. Taxonomic remarks: Pyrgulidae 

are maintained as a family although molecular data hint at a 

subfamily level (Szarowska et al., 2005). The thiarid Tinnyea 

(Hantken, 1887) is counted to the Melanopsidae. 

4.1.1. Rzehakia Lake System (RLS, ~17.5–17.2 Ma) 

In total, 39 gastropod species attributed to 15 genera (species/ 

genera ratio=2.6) have been described from the RLS in the 

papers of Rzehak (1893), Schlickum (1963, 1964a,b, 1966, 

1967), Ctyroky (1972), Steininger (1973) and Kowalke and 

Reichenbacher (2005). The fauna is characterised by its small 

size, the individuals usually being less than 10 mm in height. 

Few exceptions, such as Viviparus suevicus (Wenz, 1919) or 

Melanopsis impressa (Krauss, 1852), exceed this limit. Similarly, 

the dreissenids are small-sized with ranges from 10–20 mm 

(Harzhauser and Mandic, 2008). The highest percentage of 56% 

is contributed by Hydrobiidae (22 species). These are represented 

by Nematuerella (Sandberger, 1874) (9), Staliopsis (Rzehak, 

1893) (6) and Hydrobia (Hartmann, 1821) (2) and the endemic 

genus Ctyrokia (Schlickum, 1965) (5). All other families are 

represented by 5 or less species: Planorbidae (5), Neritidae (4), 

Melanopsidae (3), Viviparidae (2), Lymnaeidae (2), Bithyniidae 

(1). The most striking feature of the RLS gastropod fauna is the 

diversity of the hydrobiids Ctyrokia and Staliopsis. The endemicity 

on the species-level is high (77%). 

4.1.2. Dinarid Lake System (DLS, ~17–15 Ma) 

The entire gastropod fauna of the DLS is composed of 110 

species and 28 genera (species/genera ratio=3.9). The DLS 

literature is manifold, partly hard to get and a synopsis is 

completely missing up to now. The most important papers are: 

Brusina (1870, 1874, 1878, 1884b, 1881, 1896, 1897, 1902a), 

Neumayr (1869, 1880), Kittl (1895), Kochansky-Devidé and 

Slišković (1972), Jurišić-Polšak (1979), Jurišić-Polšak and 

Slišković (1988) and Olujić (1999). 

Most of the species are small-sized (b1 cm); larger shells (1– 

3 cm) are confined to few species belonging to the genus Melanopsis 

and to the thiarid Tinnyea (up to 7 cm). Hydrobiidae (40) 

and Melanopsidae (34) are the dominant families, followed by the 

Planorbidae (13). Stenothyridae (6), Pyrgulidae (6), Neritidae (5), 

Lymnaeidae (3), Viviparidae (1), Bithyniidae (1) and Valvatidae 

(1) are subordinate as taxa but may be important constituents 

concerning individual numbers. The most eye-catching radiations 

are represented by the genera Melanopsis (29), Prososthenia 

(Neumayr, 1869) (18), and Fossarulus (Neumayr, 1869) (14), 

which develop extraordinary numbers of species. All other genera 

are recorded only with 5 to 1 species. Endemic DLS genera are the 

stenothyrid Bania (Brusina, 1896), the pyrgulid Marticia 

(Brusina, 1897) and the derived clivunellids Clivunella and Delminiella, 

which are endemic even on the family level. Fossarulus 

and Dianella? Gude, 1913, although recorded as rare elements 

from other Miocene lake systems as well, display a unique 

diversity in the DLS. The endemicity level is extremely high 

(98%). 

4.1.3. Lake Skopje (LSK, ~15 Ma) 

The gastropod fauna of Lake Skopje has been described in only 

few papers (Burgerstein, 1877; Pavlovic, 1903). Sixteen smallsized 

species from 5 genera are known (species/genera ratio=3.2). 

These represent an unusual diversity of Pyrgulidae (7 species 

of Dianella) accompanied by Melanopsidae (3), Hydrobiidae 

(3), Neritidae (2) and Stenothyridae (1). There is no endemism on 

the genus level but a complete endemism on the species-level. The 

fauna is small-sized, ranging between 2 and 15 mm. 

4.1.4. Paratethyan Sarmatian Lakes (PSL, 12.4–11.8 Ma) 

The Sarmatian wetland systems were inhabited by 61 gastropod 

species attributed to 21 genera (species/genera ratio=2.9) (Hörnes, 

1856; Stoliczka, 1862; Jekelius, 1944; Boda, 1959; Harzhauser and 

Kowalke, 2002). The dominant families are the Hydrobiidae 

(15 species) and the Pyrgulidae (12) followed by the Neritidae 

(9), Valvatidae (8) and Stenothyridae (7). The rest is contributed by


Fig. 4. Family-level analysis of the gastropod faunas (note that Pyrgulidae are treated as a family and that the thiarid Tinnyea is counted to the Melanopsidae). 

Percentages are based on species numbers and do not reflect dominance of single species in individual numbers. The corresponding species numbers are given in the 

text. 

Melanopsidae (4), Planorbidae (4) and Lymnaeidae (2). Endemics 

on the genus level mainly involve stenothyrids (Aluta Jekelius, 

1932, Staja Brusina, 1897) and the pyrgulids (Socenia Jekelius, 

1944, Baglivia Brusina, 1892). All these genera persist into the Late 

Miocene and are constituents of the fauna of Lake Pannon. This 

faunistic relation to Lake Pannon is also present on the species-level 

425


and lowers the endemicity to 62%. The size of most gastropod 

species is rather small, ranging between 2 and 6 mm. Only the 

melanopsids and lymnaeids exceed the 2 cm limit. 

Within the investigated time-interval, the circum-Paratethyan 

lakes gave rise to the first peaks of diversity within the 

valvatids and neritids of the European faunas. 

4.1.5. South German lakes (incl. Lake Steinheim) (SGL, 14.3 to 

~12 Ma) 

The various small Middle Miocene lakes in southern 

Germany, including the famous Steinheim Lake, yield about 

41 gastropod species of 18 genera (species/genera ratio=2.3) 

(Klein, 1846, Gottschick, 1911, 1853, 1920; Gottschick and 

Wenz, 1916; Schlickum, 1976; Nützel and Bandel, 1993; 

Finger, 1997 and references therein). 

The Steinheim Lake experienced an outstanding endemic 

evolution of small-sized planorbids (Hilgendorf, 1867; Nützel 

and Bandel, 1993). Thus, the diversity is completely dominated 

by the Planorbidae, which are recorded with at least 24 species. 

Of these, at least 17 species belong to Gyraulus (Charpentier, 

1837). Lymnaeidae, with 7 species, are the second most speciesrich 

group, whilst the Neritidae, Melanopsidae, Bithyniidae, 

Stenothyridae, Hydrobiidae and Valvatidae are represented by 

1–3 species only. No endemism on the generic level has been 

observed so far for these lakes; a considerably endemism of 

c. 75% is represented by the species of the stenothyrids, hydrobiids 

and even more so within the planorbids. At least 10 

species are also recorded from other freshwater systems such as 

the RLS and the PSL. Aside from Tinnyea and few lymnaeids, 

the fauna is small-sized (b5 mm). 

4.1.6. Lake Pannon (LP, 11.6–5.8 Ma) 

Aside from numerous monographs treating regional Lake 

Pannon assemblages, only Müller et al. (1999) provided a familylevel 

synopsis of the mollusc fauna. The long history of Lake 

Pannon, spanning more than 6 Ma, and the large geographic 

extent of its deposits, is reflected in an enormous bibliography. In 

total, at least 497 species- and subspecies-level gastropods have 

been described from Lake Pannon. Only few of these species 

existed throughout the history of the lake. Therefore, the faunas of 

the various localities are united into three stratigraphic groups: 

LP, Phase I: 11.6–10.0 Ma: 150 gastropod species of 34 

genera are described from this early phase of Lake Pannon 

(species/genera ratio=4.4) (Brusina; 1884a, 1896, 1897, 1902a; 

Lörenthey, 1894; Halaváts, 1903; Jekelius, 1944; Papp, 1951, 

1953; 1985b, Sauerzopf, 1953). 

The early gastropod fauna of Lake Pannon is dominated by 

Melanopsidae (38), Pyrgulidae (33) and Planorbidae (29). 

Neritidae (18), Stenothyridae (11), Valvatidae (9), Hydrobiidae 

(7), Lymnaeidae (3) and Bithyniidae (1) contribute to a lesser 

amount. The endemicity is high (86%), although some species 

appeared already in the Paratethyan Sarmatian lakes. The larger 

part of the fauna is small-sized, ranging from 2–8 mm; a size 

class between 10–30 mm is frequently represented within the 

lymnaeids, planorbids and melanopsids. Nevertheless, the early 

Lake Pannon witnesses also a tendency to gigantism by species 

of the Melanopsis fossilis-complex, which may attain a 

maximum size of 80 mm, and by Tinnyea escheri vasarhelyii 

(Hantken, 1887), which may exceed 100 mm in length. 

LP, Phase II: 10.0–8.0 Ma: During the phase of the maximum 

extent of Lake Pannon, 254 species of 40 genera (species/genera 

ratio=6.4) existed (Stoliczka, 1862; Fuchs, 1873; Brusina 1884a, 

1892, 1897, 1902a; Handmann, 1882; Halaváts, 1887, 1892, 

1903, 1910; Gorjanović-Kramberger, 1901; Lörenthey, 1902; 

Moos, 1944; Papp, 1951, 1985a,b; Sauerzopf, 1953; Bartha, 

1956, 1953, 1959; Lupu, 1963; Lueger, 1979, 1980; Jiriček, 1985; 

Fordinál, 1997, 1999; Harzhauser et al., 2002). 

The increase in species richness compared to LP I is largely 

due to the radiation of Melanopsidae (60), Pyrgulidae (58), 

Planorbidae (39), Hydrobiidae (25) and Valvatidae (20). A 

slight increase in numbers is also evident for the Neritidae (23), 

Stenothyridae (14) and Lymnaeidae (10); only the Bithyniidae 

(1) remain on a low level. The species-level endemicity (89%) is 

comparable to the early Lake Pannon fauna. Moreover, the size 

structures of the faunas are comparable. 

LP, Phase III: 8.00–5.8 Ma: The late phase of Lake Pannon 

gave rise to a huge diversity of 284 gastropod species of 48 

genera (species/genera ratio=5.9) (Rolle, 1861; Fuchs, 1870a,b; 

1873; Hoernes, 1875; Herbich and Neumayr, 1875; Lörenthey, 

1893a,b,c, 1895; Brusina 1896, 1902a; Halaváts, 1887, 1897, 

1892, 1904, 1915, 1923; Gorjanović-Kramberger, 1901; Soós, 

1934; Moos, 1944; Strausz, 1951; Papp, 1951, 1985b; Sauerzopf, 

1953; Bartha, 1954; Bartha and Soós, 1955; Gillet and Marinescu, 

1971; Marinescu, 1973; Schlickum, 1978, 1953, 1979; Korpás- 

Hódi, 1983; Stevanović and Papp, 1985; Stevanović, 1941, 1978, 

1985, 1990a,b,c; Basch, 1990; Müller and Szónoky, 1990; 

Fordinál, 1994, 1996, 1998; Szilaj et al., 1999; Harzhauser and 

Binder, 2004). 

The maximum diversity of Lake Pannon III is contributed by 

Planorbidae (53), Pyrgulidae (48), Melanopsidae (41), Lymnaeidae 

(40) and Valvatidae (30). Hydrobiidae (24), Viviparidae 

(19), Neritidae (11), Stenothyridae (11) and Bithynidae (2) follow 

in decreasing numbers. Compared to LP I and LP II, an 

increase in species richness within the viviparids, planorbids and 

lymnaeids is evident, whilst the melanopsids and pyrgulids lose 

ground. The species-level endemicity remains high (83%). The 

bulk of the fauna is still represented by small-sized gastropods 

(2–8 mm), whereas the giant melanopsids have vanished. Largesized 

taxa of up to 100 mm diameter are now represented by the 

limpet-like lymnaeid Valenciennius (Rousseau, 1842). 

Although Lake Pannon is often referred to as the centre of 

Melanopsis evolution (Bandel, 2000; Geary et al., 2002), its most 

conspicuous radiations are found within the Pyrgulidae, with 

several endemic genera such as Goniochilus (Sandberger, 1875), 

Lisinskia (Brusina, 1897), Gyromelania (Wenz, 1938–1944), 

Scalimelania (Wenz, 1938–1944) and Beogradica (Pavlovic, 

1903). Microbeliscus (Sandberger, 1875), a questionable pyrgulid 

with heterostrophic protoconch, is another endemism. Among the 

Lymnaeidae, the evolution of deep-water, limpet-like morphologies 

(Provalenciennesia Gorjanović-Kramberger, 1923, Velutinopsis 

Brusina, 1884a and Valenciennius) is noteworthy. Another 

endemism is represented by the succinid Papyrotheca (Brusina, 

1893), which documents the rare adaptation of a terrestrial 

gastropod to aquatic environments.


4.1.7. Lake Dacia (LD, ~5–3 Ma) 

This initially brackish aquatic system gave rise to at least 119 

gastropod species of 21 genera (species/genera ratio=5.7) 

(Wenz, 1942; Hanganu, 1972; Hanganu and Papaianopol, 1982; 

Lubenescu and Zazuleac, 1985; Motas and Papaianopol, 1984; 

Papaianopol, 1995). The most prominent group is represented 

by the quickly radiating Viviparidae (50). Other important 

groups are the Melanopsidae (16), Hydrobiidae (15) and 

Bithyniidae (11). All other families are subordinate: Neritidae 

(9), Valvatidae (8), Planorbidae (6), Pyrgulidae (3), Lymnaeidae 

(1). Endemism is moderate on the species-level (60.5%) and 

absent on the generic level. Most species are small-sized 

(b10 mm); only the viviparids develop giant sized species of up 

to 55 mm in height. [Several elements with Lake Pannon 

affinities settled the Dacian Basin during the Late Miocene; 

herein, however, only the Pliocene assemblages are considered.] 

4.1.8. Lake Transylvania (LT, ~4.5–3 Ma) 

The fauna is mainly known from the paper of Jekelius (1932), 

who described 63 species of 17 genera from Lake Transylvania 

(species/genera ratio=3.7). The fauna is manifold and not 

dominated by a certain gastropod family. Stenothyridae (12), 

Pyrgulidae (10), Viviparidae (9) and Hydrobiidae (9) are most 

species rich, followed by Valvatidae (6), Planorbidae (6), 

Lymnaeidae (4), Melanopsidae (3), Bithyniidae (3) and Neritidae 

(1). Endemicity is high (73%) within species but absent for 

genera. The size structure of the fauna ranges from 4–17 mm and 

is rather uniform. Larger taxa are represented solely by Viviparus 

(Montfort, 1810) (b40 mm). 

4.1.9. Lake Slavonia (LS, ~4–3 Ma) 

The fauna of that lake was studied mainly during the 19th 

century. In total, 183 gastropod species of 29 genera are 

described (species/genera ratio=6.3) (Brusina, 1874, 1884b, 

1878, 1896, 1902a; Fontannes, 1886; Herbich and Neumayr, 

1875; Neumayr, 1869, 1897, 1880; Neumayr and Paul, 1875). 

The fauna is dominated by Melanopsidae (42), Viviparidae 

(35) and Hydrobiidae (28). Aside from the rare Stenothyridae 

(1) and Lymnaeidae (5), all other groups contribute in comparable 

numbers: Planorbidae (18), Neritidae (15), Valvatidae 

(16), Bithyniidae (12). The fauna is generally small-sized 

(b10 mm) except for the partly large-sized viviparids, whose 

size may exceed 50 mm. The faunistic relation of Lake Slavonia 

to Lake Pannon and Lake Transylvania is responsible for a 

moderately high endemicity of 63%. 

4.1.10. Lake Kosovo (LK, ~3–2 Ma) 

Lake Kosovo harboured a poorly diverse gastropod fauna of 

35 species of 9 genera (species/genera ratio=4; Atanacković, 

1959; Atanacković and Stevanović, 1990). Viviparidae dominate 

with 15 species, followed by the Planorbidae (12), of 

which 8 species belong to the endemic sinistral genus Kosovia. 

Other species are represented by Melanopsidae (3), Neritidae 

(2), Hydrobiidae (2) and Stenothyridae (1). The endemicity is 

very high (92%). Most of the taxa range from 10–15 mm in 

size. Larger species of up to 25 mm are represented only by 

Viviparus. 

4.1.11. Lake Šoštanj (LSO, 2.5 Ma) 

This highly endemic Late Pliocene lake fauna (endemicity 

87%) was included because of its “Miocene” fair. Only 8 species 

are reported by Rolle (1860, 1861) and Brezigar et al. 

(1985). Planorbidae and Hydrobiidae are represented by 3 and 2 

species, whilst Bithyniidae, Melanopsidae and Valvatidae are 

documented only by 1 species each. The fauna is small-sized 

(2–10 mm). Only Melanopsis and valvatids grow to 16 mm. 

4.1.12. Lake Petea (P, 0.1–0 Ma) 

Only few papers deal with the Pleistocene to Holocene 

thermal-spring lake fauna of Lake Petea. Brusina (1902b), 

Kormos (1905) and Pauca (1937) described 23 species of 9 

genera (species/genera ratio=2.6). The composition is uniquely 

dominated by Melanopsidae, whose 12 species contribute more 

than 50% to the total fauna. Planorbidae (5) are the second 

important gastropod group in Lake Petea, whilst Neritidae (3), 

Lymnaeidae (2) and Valvatidae (1) are subordinate in species 

numbers. Aside from the fully endemic melanopsid fauna, 

which raises the endemicity of the fauna to 60.8%, most taxa are 

frequently found in Pleistocene and Holocene freshwater 

systems of Europe. The fauna is very small (b8 mm) aside 

from the melanopsids (height up to 20 mm). 

5. Discussion 

5.1. Gamma diversity: size does matter but heritage is fine as 

well 

Species diversity in the studied lake systems ranges from low 

(b30; LSK, P, LSO) and moderate (30–50; RLS, SGL, LK) to 

high (51–100; PSL, LT) and very high (N100; DLS, LPI–III, 

LD). Among the classical extant long-lived lakes, only Lake 

Baikal (147 species) falls into the last grouping and even the 

high diversity class is represented only by few examples (Lake 

Tanganyika, 68; Lake Ohrid, 72) [see Brown (1994), Seddon 

(2000) and Sitnikova (1994, 2006) for data on extant lakes]. The 

high species number is correlated with high generic diversities 

(Fig. 5). This tight correlation (r 2 =0.9) is in contrast to the “gutfeeling” 

that the enormous diversities of Lake Pannon or of 

Lake Slavonia are maintained by few genera such as Melanopsis 

or Viviparus. The origin of the diversity is less easily explained. 

A simple correlation of diversity with lake size is evident on a 

very rough scale (r 2 =0.6). Thus, small systems such as Lake 

Skopje and Lake Petea yield low diversities compared to the 

huge Lake Pannon or Lake Slavonia. The relation, however, fails 

in intermediate systems. Presumably large systems such as the 

Rzehakia Lake System or the Paratethyan Sarmatian Lakes do 

not fit the pattern because they have fewer taxa than expected 

based on size. Moreover, the highest diversity is found in Lake 

Pannon III, which is smaller than Lake Pannon II. A problem of 

this approach may be the complex geometry and sometimes 

poorly known extent of the lakes, which may result in inadequate 

size estimates. 

A second ad hoc explanation for differences in γ-diversities 

is the age of the communities. A clear hint for this age/diversity 

relation is the high species richness of Lake Pannon III, which 

427


Fig. 5. A tight correlation between species-level γ-diversity and the number of 

genera exists for the analysed faunas. A similar correlation of diversity and lake 

size is evident for the endpoints but is poor for the middlefield. 

experienced more than 4 Ma of endemic evolution during the 

climatically challenging Late Miocene. This interpretation, 

however, is again too naive to be applied to all systems. Most of 

the investigated lakes with moderate to high diversities existed 

between 0.5 and b2.0 Ma. Despite the comparable ranges, the 

observed diversities differ strongly. This discrepancy is solved 

when calculating the numbers of species occurring in more than 

one lake. Early and Middle Miocene lakes have high 

endemisms and share less than 10 species. Several of the involved 

genera such as Orygoceras (Brusina, 1882) orCtyrokia 

(Schlickum, 1965) have their first appearances in these lakes 

and lack any known geological history. Starting with the 

Paratethyan Sarmatian Lakes, this pattern changes and each 

system is “vaccinated” by its ancestor (Fig. 2). About 20 species 

persist from the PSL into the faunas of the early Lake Pannon, 

contributing to an initially high diversity. Lake Pannon II 

inherited 97 species from its early stage and 88 species ultimately 

persist into the latest phase of Lake Pannon, which 

therefore displays the highest diversity observed in this study. 

The comparably short-lived Pliocene Lake Slavonia bears 28 

species which are rooted in Lake Pannon. Again, the high 

diversity is thus a matter of heritage rather than of autochthonous 

endemic evolution. The coeval Lake Dacia, after the 

take-over by freshwater settings, becomes invaded by a mixture 

of Lake Pannon species (14) and newly evolved species from 

Lake Slavonia (40). Similarly, Lake Transylvania seems to have 

profited from the evolutionary laboratory of Lake Slavonia, 

with which it shares 14 species. Thus, the first phases of radiations 

and the fastest evolutionary pulses can be postulated to 

have occurred in the Early Miocene lakes of the Balkanids 

(DLS, LSK) and in the Middle Miocene Sarmatian Paratethyan 

Lakes. The diversities of the Late Miocene and Pliocene lakes 

systems, however, were supported by gastropod lineages that 

developed in precursor-lakes. A similar mechanism was also 

documented for the extant gastropod fauna of Lake Tanganyika 

(Wilson et al., 2003). 

5.2. Phylogenetic lineages and generic inter-lake relations 

Numerous widespread non-endemic genera occur in most of 

the analysed lake faunas. Aside from the extinct thiarid Tinnyea, 

most of these genera are extant. Some extinct genera, however, 

display a striking fossil history indicating important faunal 

exchange between certain lake systems. Large stratigraphic 

gaps between the occurrences underline the highly incomplete 

record of the Neogene freshwater systems. The Dinarid Lake 

System was the presumed starting point for the evolution of the 

de-coiled planorbid Orygoceras (Brusina, 1882), the hydrobiid 

Emmericia (Brusina, 1870) and the melanopsid Melanoptychia 

(Neumayr, 1880). Thereafter, they are apparently absent from 

younger systems such as the South German Lakes and the 

Paratethyan Lakes but also from more or less coeval systems 

such as Lake Skopje and Lake Serbia. They, however, reappear 

5 Ma later in the Late Miocene as constituents of the Lake 

Pannon fauna. The DLS element Fossarulus displays an even 

larger stratigraphic gap and reappears in the Pliocene in Lake 

Kosovo. Similarly, the sinistral planorbid Kosovia appears in 

the early Middle Miocene Lake Serbia and re-enters the scene c. 

10 Ma later in Lake Kosovo. In respect to the very characteristic 

conchological features, convergent evolution is unlikely to 

be responsible for these chronologically disjunct occurrences. 

Other Dinarid Lake genera such as the hydrobiid Prososthenia 

and the pyrgulids Marticia and Dianella have a more continuous 

record and invade the coeval Lake Skopje. Afterwards, 

Prososthenia steps into the Paratethyan Sarmatian lakes, enters 

Lake Pannon and persists in Lake Transylvania, Lake Slavonia 

and Lake Dacia. Emmericia, the pyrgulid Micromelania, 

the bithyniid Tylopoma and the highly derived lymnaeid 

Valenciennius, too, manage to settle Pliocene descendants of 

Lake Pannon. Of these, only Emmericia persisted into the 

Holocene and is still found in Central Europe. 

5.3. Convergent evolution 

Aside from those partly enigmatic generic inter-lake relations, 

striking convergences help explain the numerous 

stratigraphic and biogeographic misinterpretations. Harzhauser 

and Mandic (2008) have pinpointed several examples within the 

dreissenid bivalves which developed unrelated morpho-pairs in 

the Dinarid Lake System and Lake Pannon. Even more 

astonishing is the convergent evolution of large-sized, limpetlike, 

deep-water-dwelling gastropods in these lakes. In the 

Dinarid Lake system, Clivunella and Delminiella represent 

this type. The origin of these taxa is unknown. The lymnaeid 

protoconch and earliest teleoconch of Delminiella point to 

an affiliation with the Lymnaeoidea. Clivunella lacks these 

conchological features and its ancylid early shell may point to a 

relation to the Planorboidea. These derived gastropods settled 

the deep lake habitats but were unable to spread into any other 

Early and Middle Miocene lakes. During the Late Miocene,

Lake Pannon saw a near-identical development, which led to the 

large-sized Valenciennius. In this case a good fossil record 

documents the evolution from inflated lymnaeids via various 

intermediate stages (e.g. Provalenciennesia, Vetulinopsis) to the 

depressed deep-water limpet Valenciennius. This gastropod 

managed to spread into coeval deposits of the Dacian Basin and 

survived until Pliocene times in Lake Slavonia. 

5.4. Biostratigraphy versus ecology 


Fig. 6. Cluster analysis of the total gastropod faunas of each lake system for family-, genus- and species-level (see section 2 for methods). A biostratigraphic signal is 

evident on the species-level but becomes weak at higher hierarchies. The family-level analysis reveals rather a mixture of ecology and stratigraphy. 

An analysis of the individual lake faunas on various 

taxonomic levels (family, genus, species; Fig. 6) revealed 

quite deviating patterns. The species-level cluster analysis 

clearly traces the biostratigraphic signal. Aside from the outside 

branches (Lake Skopje, Lake Šoštanj), two main clusters 

separate the Miocene and the Pliocene–Pleistocene lake faunas. 

Moreover, the Miocene cluster falls apart into an Early to 

Middle Miocene cluster and a late Middle to Late Miocene 

cluster with a distinct Lake Pannon group. This biostratigraphic 

signal begins to become lost already in the genera-based 

analysis. Again, the Miocene cluster is evident and especially 

the (PSL(LPIII(LPII LPI)))-relation is strong due to the direct 

phylogenetic and geodynamic relationship of these systems. 

The Miocene DLS fauna, however, clusters within the Pliocene 

cluster due to the contribution of freshwater genera such as 

Emmericia and Lithoglyphus. On the family level, this biostratigraphic 

grouping becomes indistinct. Instead, five distinct 

clusters are evident. Lake Petea, with the unusually high amount 

of melanopsids, represents the outgroup. The remaining cluster 

separates into planorbid-dominated lakes such as the South 

German lakes and Lake Kosovo (N37% planorbids) and a 

second cluster which divided into 3 branches: hydrobiiddominated 

lakes (DLS, RLS), pyrgulid-dominated lakes (PSL, 

LT, LP, LSK) and viviparid-dominated ones (LD, LS, LK). This 

stratigraphy-unrelated pattern reflects more ecological parameters. 

A simple salinity relation, however, is unlikely because 

the Pliocene freshwater systems Lake Transylvania and 

Lake Kosovo cluster together with the alkaline and saline 

systems such as Lake Pannon and the Paratethyan Sarmatian 

Lakes. Similarly, the hydrobiid-dominated branch unites the 

brackish Rzehakia Lake System with the freshwater Dinarid 

Lake System. Only the viviparid-dominated and the planorbid-dominated 

systems seem to correlate simply with freshwater 

settings. This is also indicated by the unionid-bivalve 

fauna of the viviparid lakes: Lake Slavonia and Lake Dacia. 

Despite the vanishing biostratigraphic signal a generalization 

is that the Early Miocene lake systems tend to be hydrobiiddominated, 

the middle-Miocene and Late Miocene are 

pyrgulid-dominated and the Pliocene systems are usually 

viviparid-dominated. 

5.5. Endemism of fossil and extant lake systems 

Most extant faunas of ancient lakes display endemism rates 

between 40 and 80% at diversities between 24 and 147 species 

(Fig. 7). The observed endemisms of the herein-considered 

fossil lake faunas is generally comparable but tend to be even 

higher (60–98%). Neither the extant nor the fossil faunas show 

any correlation between species richness and endemism. Lowdiversity 

faunas such as in Lake Skopje and Lake Kosovo 

display equally high endemism rates as the extremely diverse 

Lake Pannon and the Dinarid Lake System. The generally 

higher endemism in the fossil systems is probably related to an 

incomplete record of coeval lake faunas (e.g. Lake Skopje, Lake 

Kosovo). An exception seems to be Lake Pannon, whose 

extraordinary endemism might rather be related to adaptations 

429


Fig. 7. Endemism versus γ-diversity for fossil and extant ancient lakes [data for modern faunas from Brown (1994), Seddon (2000) and Sitnikova (1994, 2006)]. The 

endemism and species-richness of Lake Pannon, Lake Slavonia and the Dinarid Lake System are outstanding. The generally higher endemism in the fossil systems at 

comparable species numbers might hint at a still incomplete record of Neogene freshwater systems. 

of the fauna to an aberrant water chemistry coupled with a 

geological longevity. 

6. Conclusions 

Many papers dealing with extant mollusc faunas of Eurasian 

aquatic systems refer to Lake Pannon when explaining extant 

biogeographic distributions and phylogenetic relations (e.g. 

Grigorovich et al., 2003; Bunje and Lindberg, 2007). Our 

dataset, however, points to a much more complex history of the 

faunas reaching back at least to the Early Miocene. High 

endemisms and low inter-lake relations of the Early and early 

Middle Miocene lake systems suggest that these experienced 

the first autochthonous evolutionary pulses. Many genera 

display their FADs in these systems (e.g. Marticia, Kosovia, 

Orygoceras, Pyrgula, Dianella, Emmericia). This pattern 

changed at the Middle/Late Miocene boundary when Lake 

Pannon inherited numerous species which evolved prior in the 

Sarmatian Paratethyan lakes. On the generic level, parts of the 

Lake Pannon fauna can be traced back even to the Early 

Miocene faunas of the Dinarid Lake System. The combined 

effect of heritage and new radiations in a geochemically unique 

aquatic system allowed Lake Pannon to accumulate an enormous 

diversity of 497 gastropod species. Lake Pannon itself 

acted as a stepping stone for species and genera which settled 

the descendant freshwater systems such as Lake Slavonia, Lake 

Dacia and Lake Transylvania. Generic endemism thus decreased 

during the Pliocene. 

Generally, the lake faunas may be divided into pyrgulid-, 

hydrobiid-, viviparid- and planorbid-dominated lakes. The 

reason for this predominance of certain taxa is not fully understood. 

A simple relation to water chemistry is unlikely in respect 

to the similarities between the faunas of the slightly brackish 

and alkaline Lake Pannon (Harzhauser et al., 2007) and those of 

the freshwater fauna of the Dinarid Lake System. 

A weak stratigraphic signal indicates that Early Miocene 

freshwater systems are hydrobiid dominated; Middle and Late 

Miocene systems tend to be pyrgulid dominated, whilst 

Pliocene ones are often viviparid dominated. A climatic control 

is not the main force behind the pattern because the temperate 

RLS faunas are hydrobiid dominated as are the DLS faunas 

which developed during the beginning Middle Miocene 

climatic optimum. At least the switch from pyrgulid- to 

viviparid-dominated lakes in the Pliocene seems to be mainly 

explained by the pure freshwater settings that replaced the 

slightly brackish and alkaline Lake Pannon environments. 

The Neogene lake systems represent a unique laboratory of 

evolution. Examples of parallel evolution and the phenomenon 

of iterative morphologies make the analysis of ancient lake 

faunas a tantalizing endeavour. Repetitive morphologies of 

related lineages have been documented to occur in Lake 

Pannon melanopsids (Geary et al., 2002). Even more interesting 

are such iterative developments of unrelated taxa as shown 

for DLS and LP dreissenids (Harzhauser and Mandic, 2008). 

The most striking examples of such “morpho-pairs” is the DLS 

taxa Delminiella and Clivunella and the Valenciennius-lineage 

in Lake Pannon. The planorbid Clivunella and the lymnaeid 

Delminiella are two endemic limpet-like shells which, deriving 

from nearshore ancestors, adapted synchronously and independently 

to deep-water settings of the Dinarid Lake System. 

About 5 Ma later, the lymnaeids of Lake Pannon started to 

explore the deep-water habitats of that lake, resulting in the 

limpet-like Valenciennius. Such morpho-pairs have been the 

reason for frequent stratigraphic and biogeographic misinterpretations 

in the literature. Despite the huge dataset, comprising 

about 1184 gastropod taxa from 119 localities, the Neogene 

freshwater record is still poor. This fragmentary fossil record 

is underlined by disjunct stratigraphic occurrences of 

highly derived genera such as Orygoceras or Kosovia with 

gaps of 5–10 Ma.

Acknowledgements 

The studies were supported by FWF-grant P18519-B17 

“Mollusc Evolution of the Neogene Dinaride Lake System” and 

contributed to the NECLIME project. 

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Neogene lake systems of Central and South-Eastern Europe ...

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